POLLEN FLORA OF PAKISTAN LVI. VALERIANACEAE

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1 Pak. J. Bot., 39(6): , POLLEN FLORA OF PAKISTAN LVI. VALERIANACEAE ANJUM PERVEEN AND MUHAMMAD QAISER Department of Botany, University of Karachi, Karachi , Pakistan. Abstract Pollen morphology of 13 species representing 2 genera of the family Valerianaceae from Pakistan has been examined by light and scanning electron microscope. Pollen grains usually radially symmetrical, isopolar, mostly prolate-spheroidal to sub-prolate, often oblate-spheroidal rarely prolate. Tricolpate sexine thicker or thinner than nexine. Tectal surface mostly spinulose rarely scabrate or echinate. On the basis of exine ornamentation 3 distinct pollen types are recognized viz., Valeriana jatamansi-type, Valeriana hardwickii- type, Valerianella dentata-type. Introduction Valerianaceae is a family of about 13 genera and more or less 400 species, chiefly confined to temperate regions with exception of Andes of South America (Mabberley, 1987). It is represented in Pakistan by 3 genera and c. 22 species (Nasir, 1976). Valerianaceae originally described by Linneaus (1753, 1754), is divided into three tribes viz., 1) Patrinieae, 2) Triplostegieae and 3) Valerianeae by Graeber (1906). Backlund (1996) suggested that the family Valerianaceae is comprised of 14 genera, distributed in two in tribe Patrinieae (Patrinia and Nardostachys), a single genus (Triplostegia) in the tribe Triplostegieae, and the 11remaining genera are grouped into five sub tribes in the Valerianeae. Plants mostly herbs or shrubs. Annual to perennial; with or without basal aggregation of leaves, when perennial, the rhizome usually strongly scented. Leaves simple or compound, exstipulate. Flowers bisexual usually clusters with bract or bracteoles. Fruit achene-like with wing or plumose. Many members of the Valerianaceae have a distinctive odor due to the presence of valerianic acid and its derivatives. Several species have medicinal properties, and root and leaf extracts are used in treating nerve complaints. A few species are used for perfumes and dyes. Some European species of Valerianella are used in salads. However, some plants contain poisons in high concentrations. Pollen morphology of the family has been examined by Ting (1949), Erdtman (1952, 1966), Wang (1960),Taranvschi et al., ( 1966). Kuprianova & Alyoshina (1978). Nowicke & Skvarla (1979), Patel & Skvarla (1979), Blackmore & Cannon (1983), Backlund & Nilsson (1997). Clarke (1978) examined pollen morphology of family Velarianaceae in relationship to taxonomy. Clarke & Jones (1981) studied pollen morphology of the family Valerianaceae from North Western Europe. There are no reports on pollen morphology of the family Valerianaceae from Pakistan. Present investigations are based on the pollen morphology of 13 species representing 2 genera of the family Valerianaceae by light and scanning electron microscope. Materials and Methods Polleniferous material was obtained from Karachi University Herbarium (KUH) or fresh material collected from the field. The list of voucher specimens is deposited in KUH. The pollen grains were prepared for light (LM) and scanning microscopy (SEM)

2 1890 ANJUM PERVEEN & MUHAMMAD QAISER by the standard methods described by Erdtman (1952). For light microscopy, the pollen grains were mounted in unstained glycerin jelly and observations were made with a Nikon Type-2 microscope under (E40, 0.65) and oil immersion (E100, 1.25), using 10x eye piece. For SEM studies, pollen grains suspended in a drop of water were directly transferred with a fine pipette to a metallic stub using double sided cello tape and coated with gold in a sputtering chamber (Ion-sputter JFC-1100). Coating was restricted to 150A. The S.E.M examination was carried out on a Jeol microscope JSM-2. The measurements are based on readings from each specimen. Pollen diameter, polar axis (P) and equatorial diameter (E), aperture size and exine thickness were measured (Tables 1). The terminology used is in accordance with Erdtman (1952), Kremp (1965), Faegri & Iversen (1964) and Walker & Doyle (1975). General pollen characters of the family Valerianaceae Pollen grains usually radially symmetrical, isopolar. Shape mostly prolatespheroidal, to sub-prolate rarely oblate-spheroidal. Tricolpate, sexine thicker or thinner than nexine. Tectal mostly spinulose rarely echinate or scabrate. On the basis of exine ornamentation 3 distinct pollen types are recognized viz., Valeriana jatamansi-type, Valeriana hardwickii-type and Valerianella dentata-type. Key to the pollen type 1 + Tectum scabrate..... Valerianella dentata-type - Tectum not as above Tectum spinulose.. Valeriana hardwickii-type - Tectum echinate Valeriana jatamansi type. 4 Pollen type: Valeriana hardwickii-type (Fig. 1 A-F). Pollen class: 3-colpate. P/E ratio: Shape: Prolate-spheroidal to sub-prolate often oblate-spheroidal. Apertures: Colpus long narrow with acute ends. Exine: Sexine thinner than nexine. Ornamentation: Spinulose Measurements: Size: Length = (28.72-) ± 0.52 (-61.03) μm, and breadth (25.01) ± 0.59 (40.74) μm, colpi ( ) ± 0.42 (50.26) μm in long. Mesocolpium μm. Apocolpium 7.2 (8.12 ± 0.16) μm. Exine 2.5 (3.55 ± 0.12) 4.51 μm thick, sexine thicker than nexine. Tectum spinulose. Species included: Valeriana clarkei Briq., Valeriana hardwickii Wall., Valeriana pyrolifolia Decne., Valeriana himalayana Grub., Valeriana fedtschenkoi Coincy, Valeriana jaeschkeyi C.B. Clarke, Valeriana ficariifolia Boiss., Valeriana stracheyi C.B. Clarke, Valerianella muricata (Stev.) Baxt., Valerianella oxyrrhyncha Fisch., Valerianella szovitsiana Fish & May.

3 POLLEN FLORA OF PAKISTAN LVI. VALERIANACEAE 1891

1892 w A ANJUM PERVEEN & MUHAMMAD QAISER B C D E

4 1892 w A ANJUM PERVEEN & MUHAMMAD QAISER B C D E F Fig. 1. Scanning Electron micrographs of pollen grains. Valeriana fedtschenkoi: A, Polar view; B, Equatorial view; C, Exine pattern. V. pyrolifolia: D, Polar view. Valerianella muricata: E, Polar view; F, Equatorial view. Scale bar = A-F= 10 μm

Valeriana jatsmania: A, Exine pattern; B,

5 POLLEN FLORA OF PAKISTAN LVI. VALERIANACEAE 1893 A B C D E Fig. 2. Scanning Electron micrographs of pollen grains. Valeriana jatsmania: A, Exine pattern; B, Polar view; C, Equatorial view. Valerianella dentata: D, Polar view; E, Exine pattern. Scale bar = F= 10 μm

6 1894 ANJUM PERVEEN & MUHAMMAD QAISER Key to the species and groups 1 + Tectum prolate..... Valerianella szovitsiana - Tectum prolate-spheroidal to sub-prolate Pollen grains sub-prolate.. Valeriana clarkei-group (Valeriana clarkei Briq., Valeriana hardwickii Wall., Valeriana fedtschenkoi Coincy, Valeriana jaeschkeyi C.B. Clarke) - Pollen grains prolate-spheroidal... Valeriana himalayana-group (Valeriana pyrolifolia Decne., Valeriana himalayana Grub., Valeriana ficariifolia Boiss., Valeriana stracheyi C.B. Clarke, Valerianella muricata (Stev.) Baxt., Valerianella oxyrrhyncha Fisch.) Pollen type: Valeriana jatamansi-type (Fig. Fig. 2 A-C). Pollen class: 3-colpate. P/E ratio: Shape: Oblate-spheroidal. Apertures: Colpi long rounded acute ends. Exine: Sexine thicker than nexine. Ornamentation: Echinate, echines with cushions. Measurements: Size: Length = (37.5-) ± 0.52 (-45.8) μm, and breadth (35.01) ± 0.59 (54.74) μm, colpi ( ) ± 0.42 (32.5) μm in long. Mesocolpium μm. Apocolpium 7.5 (8.12± 0.16) 8.75 μm. Exine 2.5 (3.2 1 ± 0.11) 4.0 μm thick, sexine thicker than nexine. Tectum echinate, echines with cushions. Species included: Valeriana jatamansi Jones Pollen type: Valerianella dentata-type (Fig. 2 D & E). Pollen class: 3-colpate. P/E ratio: Shape: Oblate-spheroidal. Apertures: Colpus long narrow with acute ends. Exine: Sexine thicker than nexine. Ornamentation: Scabrate. Measurements: Size: Length = (30.5-) 33.8 ± 0.2 (-37.8) μm and breadth (31.25) 33.5 ± 0.6 (36.25) μm, colpi (18.75-) 21.5 ± 0.76 (25.8) μm long. Mesocolpium μm. Apocolpium (6.23-) 6.50 ± 0.61 (7.5) μm. Exine 2.5 (3.0 ± 0.5) 3.5 μm thick, sexine thicker than nexine. Tectum scabrate-punctate. Species included: Valerianella dentata (L.) Poll. Discussion Family Valerianaceae is more or less stenopalynous family. Pollen grains generally prolate-spheroidal, rarely oblate-spheroidalor prolate. Tricolporate with spinulose Tectum. However, little variation in exine pattern has been observed. On the basis of exine ornamentation family is divided into three pollen types. Viz., Valeriana jatamansitype Valeriana hardwickii-type and Valerianella dentata-type. Backlund & Nilsson (1997) reported the spheroidal, tectate, triporate to tricolpate pollen grains in the families Caprifoliaceae, Dipsacaceae and Valerianaceae.

7 POLLEN FLORA OF PAKISTAN LVI. VALERIANACEAE 1895 Number of workers like Takhtajan (1980), Cronquist (1981), Throne (1983) and Dahlgren (1989) kept the family Valerianaceae under the order Dipsacales near Dipsacaceae. Molecular data including sequence data from the chloroplast genes rbcl (Donoghue et al., 1992) and ndhf (Pyck et al., 1999), restriction site data (Downie & Palmer, 1992), support the placement of Valerianaeae as a sister group to Dipsacaceae. Both of these herbaceous groups are united by having distinctive pollen morphology and chlorophyllous embryos, a trait unique within Dipsacales (Backlund and Donoghue, 1996). The close relationship between Valerianaceae and Dipsacaceae is further supported by other morphological characters also like, simple perforated vessel modification of calyx lobes, and reduced endosperm (Judd et al., 1994; Manchester & Donoghue, 1995). Two major lineages have been suggested by Clarke (1978), the Valeriana group, with large spines or verrucae and simple, unbranched, and uniform columellae; and the Valerianella group, with small to medium-sized spines and branched columellae increasingly longer and more clearly branched below the poles. Van der Hammen (1979), reported Valeriana pollen among the elements of the primitive paramo flora originating in the upper Pliocene and lower Pleistocene. References Backlund, A In A. Backlund: Phylogeny of the Dipsacales, Doctoral thesis in systematic botany from Uppsala University, Sweden. Backlund, A. and A. Nilsson Pollen morphology and the systematic position of Triplostegia (Dipsacales). Taxon, 46: Blackmore, S. and M.J. Cannon Palynology and systematics of Morinaceae. Rev. Palaeobot. Palynol., 40: Clarke, G Pollen morphology and generic relationships in the Valerianaceae. Grana., 17: Clarke, G The northwest European pollen flora Dipsacaceae. Rev. Palaeobot. Palynol., 33: Clarke, G. and M.R. Jones The northwest European pollen flora Valerianaceae. Rev. Palaeobot. Palynol., 24: Cronquist, A An Integrated System of Classification of Flowering Plants. Columbia Univ. Press, New York. Dahlgren, G The last Dahlgrenogram. In: System of classification of dicotyledons. (Ed.): Kit.-Tan. The Davis and Hedge Fetschrift. Edinburgh Univ., Press. Donoghue, M.J., R.G. Olmstead, J.F. Smith and J.D. Palmer Phylogenetic relationships of Dipsacales based on rbcl sequences. Annals of the Missouri Botanic Gardens, 79: Downie, S.R. and J.D. Palmer Restriction site mapping of the chloroplast DNA inverted repeat: a molecular phylogeny of the Asteridae. Annals of the Missouri Botanic Gardens, 79: Erdtman, G Pollen Morphology and Plant Taxonomy. Angiosperms. Chronica Botanica Co., Waltham, Massachusettes. Erdtman, G Pollen morphology and plant taxonomy: angiosperms. Hafner, New York. Faegri, K. and J. Iversen Text book of Pollen Analysis. Munksgaard, Graeber, P Valerianaceae andinae. Bot. Jahrb. Syst. 37: Judd, W.S., R.W. Sanders and M.J. Donoghue Angiosperm family pairs: preliminary phylogenetic analyses. Harvard Papers Bot., 5: pp Kremp, G.O.W Encyclopaedia of Pollen Morphology, Univ. Arizona Press, Tuscon, U.S.A. Kuprianova, L.A. and L.A. Alyoshina Pollen dicotyledoneaerum Florae Partis Europareae. URSS. Lamiaceae-Zygophyllaceae. (in Russian). Nauka 184 p. Akad. Sci. USSR.L. Komarov. Inst. Bot.

8 1896 ANJUM PERVEEN & MUHAMMAD QAISER Linnaeus, C Species plantarum. Stockholm. Linnaeus, C Genera plantarum. Stockholm. Mabberley, D.J The Plant Book, a Portable Dictionary of Higher Plants. Cambridge University Press, Cambridge. Manchester, S.R. and M.J. Donoghue Winged fruits of Linnaeeae (Caprifoliaceae) in the Tertiary of Western North America: Diplodipelta gen. nov. International Journal of Plant Science, 156: Nasir, Y Valerianaceae. In: Flora of Pakistan. (Eds.): S.I. Ali & E. Nasir. 101: Nowicke, J.W. and J.J. Skvarla Pollen morphology: The potential influence in higher order systematics. Ann. Mo. Bot. Gard., 66: Patel, V.C. and J.J. Skvarla Valerianaceae pollen morphology, Pollen & Spores 21: Pyck, N., P. Roels and E. Smets Tribal relationships in Caprifoliaceae: evidence from a cladistic analysis using ndhf sequences. Syst. Geogr. Pl., 69: Takhtajan, A Outline of the classification of flowering plants (Magnoliophyta). Bot. Rev. (Lancaster), 46: Tarnavschi, I.T., N. Mitroiu and D. Radulescu Studiul palinologic al representantilor ordinuluri Rubiales. An. Univ. Buc. Ser. Biol., 15: Thorne, R.F Proposed new realignments in the angiosperms. Nordic J. Bot., 3: 117. Ting, S.U Illustration of pollen grains of some Chinese plants. Bot. Notiser, 4: Van der Hammen The Pleistocene changes of vegetation and climate in tropical South America. Journal of Biogeography, 1: Walker, J.W. and J.A. Doyle The basis of Angiosperm phylogeny: Palynology. Ann. Mo. Bot. Gard., 62: Wang, F.H Pollen grains of China. (in Chinese). 276 p. (Received for publication 8 May 2007)

POLLEN FLORA OF PAKISTAN XLI. CUSCUTACEAE

Pak. J. Bot., 36(3): 475480, 2004. POLLEN FLORA OF PAKISTAN XLI. CUSCUTACEAE ANJUM PERVEEN AND MOHAMMAD QAISER Department of Botany, University of Karachi, Karachi, Pakistan Abstract Pollen morphology